Neanderthal behavior is subject to much study and speculation. Neanderthals were thought to be almost exclusively carnivorous[1] and apex predators.[2] However, new studies indicate that they had cooked vegetables in their diet.[3][4] They made advanced tools, had language (the nature of which is debated) and lived in complex social groups.

Neanderthal brains were somewhat larger than humans but were shaped a bit differently[5] since they evolved separately for several hundred thousand years. The size and distribution of archaeological sites shows that Neanderthals spent their lives mostly in small groups of 5-10 individuals (compared to 20-30 individuals for Cro-Magnon man). Elders were rare as few Neanderthals lived past 35.

Skeletal evidence shows that injured individuals were often nursed back to health by others. Neanderthals rarely made contact with outsiders or traveled outside their small home territories. Although many Neanderthal sites have rare pieces of high-quality stone from more than 100 kilometers away, there is not enough to indicate trade or even regular contact with other communities.

It has been suggested by one pair of researchers that these stones may instead be "gifts" brought by adolescents wishing to join a new community (some form of "marrying out" was essential due to the small size of Neanderthal territories). In their view, this lack of trade could indicate that Neanderthals may have lacked some cognitive abilities for dealing with strangers, such as "cheater detection" and the ability to judge the value of one commodity in terms of another. Neanderthals had a smaller cognitive part of the brain and this would have limited them, including their ability to form larger groups.[6] The quality of tools found at archaeological sites is further said to suggest that Neanderthals were good at "expert" cognition, a form of observational learning and practice acquired through apprenticeship that relies heavily on long-term procedural memory.[7]

Neanderthal toolmaking supposedly changed little over hundreds of thousands of years. The lack of innovation was said to imply they may have had a reduced capacity for thinking by analogy and less working memory. The researchers further speculated that Neanderthal behavior would probably seem neophobic, dogmatic and xenophobic to modern humans.[7]

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The idea that Neanderthals lacked complex language was once widespread,[8] despite concerns about the accuracy of reconstructions of the Neanderthal vocal tract, until 1983, when a Neanderthal hyoid bone was found at the Kebara Cave in Israel. The hyoid is a small bone that connects the musculature of the tongue and the larynx, and by bracing these structures against each other, allows a wider range of tongue and laryngeal movements than would otherwise have been possible. The presence of this bone implies that structured speech was anatomically possible and that the repertory of sounds that might be produced was wide enough to contain well-defined sets of phonemes, and not simply inarticulate guttural grunts. The bone found is virtually identical to that of modern humans.[9] A recent study on the Kebara hyoid has used X-ray microtomography and finite element analysis to compare it to modern human hyoids. The study concluded that the Neanderthal hyoid showed histological features and micro-chemical behavior similar to a modern human's hyoid, indicating that the two were used in the same way.[10]

There is some circumstantial evidence for thinking that Neanderthals had language with words and some kind of syntax; some of their tool-making and hunting tactics would have been difficult to learn and execute without it.[11] A recent extraction of DNA from Neanderthal bones indicates that Neanderthals had the same version of the FOXP2 gene as modern humans. This gene is known to play a role in human language.[12]

The morphology of the outer and middle ear of Neanderthal ancestors, Homo heidelbergensis, found in Spain, suggests they had an auditory sensitivity similar to modern humans and very different from chimpanzees. They were probably able to differentiate between many different sounds.[13]

Neurological evidence for potential speech in neanderthalensis may exist in the form of the hypoglossal canal. The canal of neanderthalensis is the same size or larger than in modern humans, which is significantly larger than the canal of australopithecines and modern chimpanzees. The canal carries the hypoglossal nerve, which controls the muscles of the tongue. This suggests to some theorists that neanderthalensis had vocal capabilities similar to modern humans.[14]

A research team from the University of California, Berkeley, led by David DeGusta, however, argues that the size of the hypoglossal canal is not an indicator of speech. His team's research, which shows no correlation between canal size and speech potential, shows there are a number of extant non-human primates and fossilized australopithecines that have an equal or larger hypoglossal canal.[15]

Another anatomical difference between Neanderthals and modern humans is the former's lack of a mental protuberance (the point at the tip of the chin). This may be relevant to speech, as the mentalis muscle contributes to moving the lower lip and is used to articulate a bilabial click. While some Neanderthal individuals do possess a mental protuberance, their chins never show the inverted T-shape of modern humans.[16] In contrast, some Neanderthal individuals show inferior lateral mental tubercles (small bumps at the side of the chin).

Steven Mithen (2006) speculates that the Neanderthals may have had an elaborate proto-linguistic system of communication that was more musical than modern human language, and that pre-dated the separation of language and music into two separate modes of cognition. He called this hypothetical lingual system 'hmmmmm' because it would be Holistic, manipulative, multi-modal, musical and mimetic.[17]

Neanderthal and Middle Paleolithic archaeological sites show a smaller and different toolkit than those found in Upper Paleolithic sites, which were perhaps occupied by modern humans that superseded them. Fossil evidence indicating who may have made the tools found in Early Upper Paleolithic sites is still missing. A survey of 332 archeological sites occupied over a period of 200,000 years under varying climatic conditions using lithic tool data from 190 layers at 103 sites showed that the Neanderthal toolkit changed little, showing technological inertia, a slower rate of variability compared to modern humans whose toolkits show more economic reactivity, variety in response to changing conditions. In addition to the obvious[to whom?] hypothesis that Neanderthals were not very creative despite having larger brains than modern humans, an alternate demographic hypothesis is that there were never very many Neanderthals, perhaps less than 10,000, making the probability of innovation low.[18]

An artist's impression of Neanderthal life.

Neanderthals used tools of the Mousterian class, which were often produced using soft hammer percussion, with hammers made of materials like bones, antlers, and wood, rather than hard hammer percussion, using stone hammers. A result of this is that their bone industry was relatively simple. However, there is good evidence that they routinely constructed a variety of stone implements. Neanderthal (Mousterian) tools most often consisted of sophisticated stone-flakes, task-specific hand axes, and spears. Many of these tools were very sharp. There is also good evidence that they used a lot of wood, but wooden objects are unlikely to have survived to the present.[19]

Neanderthals were capable of building dugout boats since the Middle Paleolithic.[20][21] Mousterian stone tools discovered on the southern Ionian Greek islands suggests that Neanderthals were navigating the Mediterranean Sea as early as 110,000 years BP.[22][23] Quartz hand-axes, three-sided picks, and stone cleavers from Crete have also been recovered that date back about 170,000 years BP.[24]

There is some evidence for interpersonal violence among Neanderthals. A 36,000-year-old Neanderthal skull found near St. Césaire has a healed fracture in its cranial vault that was most likely caused by the impact of a sharp implement. The location of the wound suggests interpersonal violence rather than an accident. Because the wound healed, it is known that the individual survived the attack.[25]

Also, while they had weapons, whether they had projectile weapons is controversial. They had spears, made of long wooden shafts with spearheads firmly attached, but some think these were thrusting spears and not projectiles.[26] Still, a Levallois point embedded in an animal vertebra shows an angle of impact suggesting that it entered by a "parabolic trajectory" suggesting that it was the tip of a projectile.[27]

Moreover, a number of 400,000-year-old wooden projectile spears were found at Schöningen in northern Germany. These are thought to have been made by the Neanderthals’ ancestors, Homo erectus or Homo heidelbergensis. Generally, projectile weapons are more commonly associated with H. sapiens. The lack of projectile weaponry may be an indication of different sustenance methods, rather than inferior technology or abilities. The situation is identical to that of native New Zealand Māori—modern Homo sapiens, who also rarely threw objects, but used spears and clubs instead.[28] Neanderthals rarely produced bone tools from their prey. Neanderthals apparently did not have needles, but at best, bone awls to drill eyelets for lacing skins and furs together.[29] Some tools may have been due to trade or copying from Homo sapiens who coexisted with Neanderthals near the end of the latter's existence.[30]

Neanderthals also performed many sophisticated tasks normally associated only with modern humans. For example, they controlled fire, constructed complex shelters, and skinned animals. A trap excavated at La Cotte de St Brelade in Jersey gives testament to their intelligence and success as hunters.[31]

Particularly intriguing is a hollowed-out bear femur with holes that may have been deliberately bored into it, known as the Divje Babe flute. This bone was found in western Slovenia in 1995, near a Mousterian fireplace, but its significance is still disputed. Some paleoanthropologists hypothesize it was a musical instrument, others believe it was not the work of Neanderthals, or that the chomping action of another bear made the holes.

Pendants and other jewelry showing traces of ochre dye and of deliberate grooving have also been found[32] with later finds, particularly in France, but whether they were created by Neanderthals or traded to them by Cro-Magnons is a matter of controversy.

Although much has been made of the Neanderthals' burial of their dead, their burials were less elaborate than those of anatomically modern humans. The interpretation of the skeleton known as Shanidar IV as having been buried with flowers, and therefore being a form of ritual burial,[33] has been questioned.[34]Paul Pettitt has stated that the "deliberate placement of flowers has now been convincingly eliminated", noting that "A recent examination of the microfauna from the strata into which the grave was cut suggests that the pollen was deposited by the burrowing rodent Meriones tersicus, which is common in the Shanidar microfauna and whose burrowing activity can be observed today".[35]

Early studies indicated that Neanderthals were highly carnivorous and obtained most of the protein in their diet from animal sources.[36] Recently, however, traces of fossilized plants have been extracted from Neanderthal teeth found in Belgium, Iraq, and East Africa indicating they also ate plants such as grains and legumes in addition to meat.[37]

Neanderthals hunted large animals, such as the mammoth. Stone-tipped wooden spears were used for hunting and stone knives and poleaxes were used for butchering the animals or as weapons. However, they are believed to have practiced cannibalism or ritual defleshing. This hypothesis was formulated after researchers found marks on Neanderthal bones similar to the bones of a dead deer butchered by Neanderthals.

Intentional burial and the inclusion of grave goods are the most typical representations of ritual behavior in the Neanderthals and denote a developing ideology. However, another much debated and controversial manifestation of this ritual treatment of the dead comes from the evidence of cut-marks on the bone, which has 'historically been viewed' as evidence of ritual defleshing.

Neanderthal bones from various sites (Combe-Grenal and Abri Moula in France, Krapina in Croatia and Grotta Guattari in Italy) have all been cited as bearing cut marks made by stone tools.[38] However, the results of technological tests have revealed varied causes.

Re-evaluation of these marks using high-powered microscopes, comparisons to contemporary butchered animal remains, and recent ethnographic cases of excarnation mortuary practises have shown that perhaps this was a case of ritual defleshing.

At Grotta Guattari, the apparently purposefully widened base of the skull (for access to the brain) has been shown to be caused by carnivore action, with hyena tooth marks found on the skull and mandible.

According to some studies, fragments of bones from Krapina show marks similar to those on bones from secondary burials at a Michigan ossuary (14th century AD), and are indicative of removing the flesh of a partially decomposed body.

According to others, the marks on the bones found at Krapina are indicative of defleshing, although whether this was for nutritional or ritual purposes cannot be determined with certainty.[39]

Evidence of cannibalism includes:

Analysis of bones from Abri Moula in France does seem to suggest cannibalism was practiced here. Cut-marks are concentrated in places expected in the case of butchery, instead of defleshing. Additionally the treatment of the bones was similar to that of roe deer bones, assumed to be food remains, found in the same shelter.[40]

At El Sidron in Northern Spain, scientists have found evidence pointing to the cannibalism of 12 individuals by what is hypothesized to have been a neighboring group of Neanderthals. According to Carles Lalueza-Fox of the Institute of Evolutionary Biology in Barcelona, the individuals (three children aged from two to nine, three teenagers, and six adults) appear to have been "killed and eaten, with their bones and skulls split open to extract the marrow, tongue and brains." Scientists believe that the lack of any evidence of a fire makes it likely that the event happened in winter, during times when food was scarce.[41]

Evidence indicating cannibalism would not distinguish Neanderthals from modern Homo sapiens. Ancient and existing Homo sapiens are known to have practiced cannibalism and/or mortuary defleshing (e.g., the sky burial of Tibet).

Grooves in bones are hypothesized to be cuts by Neanderthal tools, not animal teeth. The chances of them being random, as some writers attributing them to animals have proposed, is debated.

A 2009 report in the Proceedings of the National Academy of Sciences (PNAS) on two archaeological sites in the Murcia province of southern Spain records the discovery of shells showing pigment residues, and concludes that these were used by the Neanderthals as make-up containers. Sticks of the black pigment manganese have previously been discovered in Africa. These may have been used as body paint by Neanderthals.[42] According to one study, Neanderthal use of pigments including red ochre was restricted to interglacial periods, disappearing during glacial periods.[43]

Recent new dating techniques of cave art has suggested the possibility Neanderthals were cave painters. Many cave paintings are much older than previously thought, and possibly pre-date the arrival of H. sapiens.[44] Symbolism or simple art work has been found in 2012, inside the Gorham's Cave in Gibraltar; it may have been a sign indicating to others that the cave was already inhabited.[45]

While some Neanderthals used caves for shelter, the Molodova archaeological site in eastern Ukraine suggests others built dwellings using animal bones. A building was made of mammoth skulls, jaws, tusks and leg bones, and had 25 hearths inside.[46]